Multimodal signaling in the Small Torrent Frog (Micrixalus saxicola) in a complex acoustic environment

Many animals use multimodal (both visual and acoustic) components in courtship signals. The acoustic communication of anuran amphibians can be masked by the presence of environmental background noise, and multimodal displays may enhance receiver detection in complex acoustic environments. In the present study, we measured sound pressure levels of concurrently calling males of the Small Torrent Frog (Micrixalus saxicola) and used acoustic playbacks and an inflatable balloon mimicking a vocal sac to investigate male responses to controlled unimodal (acoustic) and multimodal (acoustic and visual) dynamic stimuli in the frogs’ natural habitat. Our results suggest that abiotic noise of the stream does not constrain signal detection, but males are faced with acoustic interference and masking from conspecific chorus noise. Multimodal stimuli elicited greater response from males and triggered significantly more visual signal responses than unimodal stimuli. We suggest that the vocal sac acts as a visual cue and improves detection and discrimination of acoustic signals by making them more salient to receivers amidst complex biotic background noise.     

Limb regeneration and subsequent asymmetry in a male secondary sexual character influences sexual selection in wolf spiders

Males of the brush-legged wolf spider Schizocosa ocreata (Hentz) have conspicuously decorated forelegs used in courtship and agonistic displays. Approximately one in five juvenile males has a missing or regenerating foreleg, and regeneration of a leg lost during development usually results in the absence of a decorative tuft on that leg. The subsequent asymmetry in this male secondary character significantly decreases success in both courtship of females and male-male agonistic interactions. Experimental removal of tufts from one leg of previously successful symmetric males produces similar results. As a test for concomitant behavioral effects, female spiders were shown video images of a courting male with symmetric tufts and the same video image altered to have asymmetric tufts. Female receptivity to the asymmetric video image was lower. In contrast to fluctuating asymmetry resulting from developmental instability, leg tuft asymmetry in S. ocreata most likely arises from a single event during ontogeny – possibly leg loss from an aggressive or predator encounter  – and may serve as a quality indicator in female mate choice. Received: 27 July 1995/ Accepted after revision: 19 November 1995     

Effect of visual background complexity and light level on the detection of visual signals of male <Emphasis Type="Italic">Schizocosa ocreata</Emphasis> wolf spiders by female conspecifics

For visual signaling to be effective, animal signals must be detected and discriminated by receivers, often against complex visual backgrounds with varying light levels. Accordingly, in many species, conspicuous visual displays and ornaments have evolved as a means to enhance background contrast and thereby increase the detection and discrimination of male courtship signals by females. Using video playbacks, we tested the hypothesis that visual courtship displays and leg decorations of male Schizocosa ocreata wolf spiders are more conspicuous against complex leaf-litter backgrounds. Video exemplars of courting males with manipulated leg tufts were superimposed on different backgrounds (complex leaf litter in sun or shade, featureless gray background) and presented to female spiders. Females were more likely to orient to males presented against lighter backgrounds (litter in sun, gray) than the darker ones (litter—shade). Males with larger tufts were also more likely to be detected, as latency to orient was shortest for enlarged and longest for removed tufts. Latency of females to approach was shorter against lighter backgrounds, and approach latency was longest for males without tufts. Female receptivity scores were significantly greater for males against lighter backgrounds, and males with larger tufts had higher scores. These results suggest that both complexity and light level of display backgrounds affect the detection of male visual courtship signals by females and that aspects of the male phenotype may increase chances of detection (and receptivity) against visually complex backgrounds.     

Sexual selection,male morphology,and the efficacy of courtship signalling in two wolf spiders (Araneae: Lycosidae)

Males of the brush-legged wolf spider, Schizocosa ocreata (Araneae: Lycosidae), possess a conspicuous male secondary sexual character: dark pigmentation and tufts of bristles on the tibiae of their forelegs. We tested several hypotheses relating to the role of this conspicuous trait in sexual selection. Triad mating experiments suggest that the tufts do not play an obvious role in the operation of sexual selection by either male competition or female choice, as there were no significant differences in the mating success of intact and experimentally shaved males. However, females mated more often with males that initiated courtship first, suggesting that capture of a female’s attention by male signalling may play a critical role. In behavioral experiments that paired a single male with a female in arenas that allowed both visual and vibratory signal transmission during courtship, female receptivity did not vary significantly with the presence or absence of tufts. However, experiments that isolated the visual component of communication (by eliminating vibratory communication) revealed a significant effect of the presence of tufts: females showed receptivity less often to males with tufts removed. Female response to visual signals was much greater in S. ocreata than in its sibling congener, Schizocosa rovneri, which lacks male tufts. We hypothesize that the tufts serve to increase the efficacy of visual displays of S. ocreata, as vibratory communication is constrained by the complex leaf litter habitat of some populations. Such environmental constraints may make visual signalling over distance a critical factor for effective courtship communication, which may in turn strongly influence male fitness. Received: 30 September 1994/Accepted after revision: 4 August 1995     

Preference for male traits in female wolf spiders varies with the choice of available males,female age and reproductive state

Individual variation in female preference for male traits may influence mate choice, especially if benefits and costs of choosiness vary with the range of available males or reproductive timing. We examined variation in female preference for male leg tuft size in Schizocosa ocreata (Hentz) wolf spiders with video playback. Dichotomous (simultaneous) choice test experiments included all possible combinations of four stimuli (i.e., modified versions of the same video male stimulus): average tuft size (control), reduced (−25%), enlarged (+25%), and no tufts (removed). Females exhibited a directional preference for larger tuft size independent of the nature of the choice (except for reduced tufts vs no tufts where no difference was seen). Female preference in the short term (over a period of 4 days) was also highly repeatable for control vs reduced tufts, but not for control vs enlarged tufts. Responses of females in ‘no-choice’ presentations of a single (control) male stimulus varied with age post-maturity; females were less receptive in weeks 1 and 2, highly receptive at week 3, and less thereafter. Mated females were least receptive and most aggressive towards a male stimulus. Females offered choices repeatedly at different ages post-maturity consistently preferred the control male vs reduced tufts over all 3 weeks but varied in their preference for enlarged tufts vs control male. In the first 2 weeks, females preferred the enlarged tuft male stimulus, but showed no preference by the third week. Females tested in week 4 showed no preference in either choice. Results suggest that the potential interaction between female preference for male traits and female reproductive timing may be a critical consideration in mate choice.     

Trends in space occupation by the encrusting sponge Crambe crambe: variation in shape as a function of size and environment

The relationship between sponge size, habitat and shape was studied in the encrusting sponge Crambe crambe (Schmidt, 1862), which is distributed widely throughout the shallow Mediterranean littoral. Examination of sponge patches in shaded and well-illuminated habitats showed that the degree of peripheral irregularity of the edges of a patch is directly related to patch size. This relationship is valid only for sponges of >100 mm² in area. Photophilic and sciaphilous sponges display different growth forms. The pattern of growth is interpreted in terms of competition for space. The directional growth of sciaphilous sponges may be due to the presence of dominant neighbours that are good space competitors, and the irregular growth of photophilic sponges to the absence of such neighbours.     

鼎湖山马尾松林凋落物分解对凋落物输入变化的响应

为了了解我国南方森林常见的人为干扰（凋落物收取）活动对生态系统养分循环的影响,研究了鼎湖山马尾松林3种主要树种凋落物分解及其养分释放对凋落物输入量变化的响应。这3种树种分别为马尾松（Pinus massoniana）、荷木（Schimasuperba）和锥栗（Castanopsis chinensis）。凋落物输入量变化分别为凋落物去除（L-）、加倍（L＋）和对照（L）3种处理,每种处理25个重复。经过18个月的处理试验,凋落物分解速率及其养分释放随树种、分解阶段和凋落物处理不同而异。荷木、马尾松和锥栗分解物平均残留率分别为0.46±0.01、0.42±0.01、0.40±0.02,其中,荷木与锥栗、马尾松差异性显著。不同处理间的凋落物分解速率差异显著,加倍、对照和去除处理样地凋落物的平均残留率分别为0.51±0.08、0.53±0.09和0.55±0.08。凋落物加倍处理促进了凋落物分解过程中C的释放,而去除凋落物处理则抑制了N、P的释放。以上结果表明,凋落物收取活动不仅直接带走凋落物中的大量养分,而且抑制了凋落物分解及其养分释放。     

Chemoreceptors distribution and relative importance of male forelegs and palps in intersexual chemical communication of the wolf spider Pardosa astrigera

The chemoreceptors distributed on forelegs and palps of male wolf spiders are known to be involved in intersexual chemical communication, yet their distribution pattern and relative importance are supposed. This study employed scanning electron microscopy (SEM) to determine chemoreceptor morphology and distribution pattern on male forelegs and palps of the wolf spider Pardosa astrigera. We also performed behavioral assays to determine their relative importance in intersexual chemical communication. Although the fine structure of the chemoreceptors distributed on male forelegs and palps did not differ greatly, the chemoreceptors were more widely distributed on palps relative to forelegs. When the chemoreceptors of palps were chemically ablated with zinc sulfate, male courtship latency was significantly prolonged and courtship intensity significantly suppressed as compared to those of the forelegs-ablated and/or control ones. Therefore, we conclude that the contact chemoreceptors are mostly distributed on male palps, thereby males locating their mates through dragline-mediated female pheromone.     

Spectral reflectance and communication in the wolf spider, <Emphasis Type="Italic">Schizocosa ocreata</Emphasis> (Hentz): simultaneous crypsis and background contrast in visual signals

Animal color patterns often reflect a compromise between natural selection for crypsis or inconspicuousness to predators and sexual selection for conspicuousness to potential mates. In leaf litter-dwelling wolf spider species like Schizocosa ocreata, body coloration often closely matches the background coloration of a generally brown environment. However, body parts used in communication should exhibit high contrast against background coloration. We used spectral analysis to examine male and female S. ocreata for matching and contrasting coloration against leaf litter. Values were plotted in multivariate color space, based on reflectivity in different frequency ranges. When viewed from above, colors of both males and females overlap with values for dead brown leaf litter and soil, suggesting cryptic coloration when viewed by potential predators. However, when viewed from a lateral perspective, both males and females show color values that are polar opposites of litter backgrounds, suggesting higher contrast when viewed by other spiders. Moreover, male secondary characters used in visual signaling by S. ocreata (tibia brushes) show the highest level of background contrast. These findings suggest that S. ocreata wolf spiders have color patterns that provide both crypsis and background contrast at the same time, depending on receiver viewing perspective.     

Patterns of litter disappearance in a northern hardwood forest invaded by exotic earthworms.

A field study was conducted to evaluate the effects of exotic earthworm invasions on the rates of leaf litter disappearance in a northern hardwood forest in southcentral New York, USA. Specifically, we assessed whether differences in litter quality and the species composition of exotic earthworm communities affected leaf litter disappearance rates. Two forest sites with contrasting communities of exotic earthworms were selected, and disappearance rates of sugar maple and red oak litter were estimated in litter boxes in adjacent earthworm-free, transition, and earthworm-invaded plots within each site. After 540 days in the field, 1.7-3 times more litter remained in the reference plots than in the earthworm-invaded plots. In the earthworm-invaded plots, rates of disappearance of sugar maple litter were higher than for oak litter during the first year, but by the end of the experiment, the amount of sugar maple and oak litter remaining in the earthworm-invaded plots was identical within each site. The composition of the earthworm communities significantly affected the patterns of litter disappearance. In the site dominated by the anecic earthworm Lumbricus terrestris and the endogeic Aporrectodea tuberculata, the percentage of litter remaining after 540 days (approximately 17%) was significantly less than at the site dominated by L. rubellus and Octolasion tyrtaeum (approximately 27%). This difference may be attributed to the differences in feeding behavior of the two litter-feeding species: L. terrestris buries entire leaves in vertical burrows, whereas L. rubellus usually feeds on litter at the soil surface, leaving behind leaf petioles and veins. Our results showed that earthworms not only accelerate litter disappearance rates, but also may reduce the differences in decomposition rates that result from different litter qualities at later stages of decay. Similarly, our results indicate that earthworm effects on decomposition vary with earthworm community composition. Furthermore, because earthworm invasion can involve a predictable shift in community structure along invasion fronts or through time, the community dynamics of invasion are important in predicting the spatial and temporal effects of earthworm invasion on litter decomposition, especially at later stages of decay.     

Evaluation of wind environment around a residential complex using a PIV velocity field measurement technique

Wind-tunnel simulations were employed to evaluate the wind environment around a tested residential area located near industrial complexes. The scaled-down geomorphological model of the test area was placed in the test section of a boundary layer wind tunnel. Particle image velocimetry (PIV) measurements were made in five vertical planes and one horizontal plane around the test area for two prevailing wind directions. The results showed that the wind speed decreased in the near surface layer and the velocity fluctuations increased in the upper region due to the presence of hills and high-rise buildings around the test area. Regions of flow separation and low-speed flow were found inside the test area for both the wind directions. The result suggests that the high-rise buildings should be well arranged with respect to the main wind directions to increase the natural ventilation inside the residential complex at the initial design stage.     

Variation in the temporal and spatial use of signals and its implications for multimodal communication

The use of signals across multiple sensory modalities in communication is common in animals and plants. Determining the information that each signal component conveys has provided unique insights into why multimodal signals evolve. However, how these complex signals are assessed by receivers will also influence their evolution, a hypothesis that has received less attention. Here, we explore multimodal signal assessment in a closely related complex of island flycatchers that have diverged in visual and acoustic signals. Using field experiments that manipulated song and plumage colour, we tested if song, a possible long-range signal, is assessed before plumage colour in conspecific recognition. We find that divergent song and colour are assessed in sequence, and this pattern of sequential assessment is likely mediated by habitat structure and the extent of differences in signal characteristics. A broad survey of the literature suggests that many organisms from a wide range of taxa sequentially assess multimodal signals, with long-range signals attracting conspecifics for further assessment of close-range signals. Our results highlight the need to consider how signals are assessed when understanding multimodal signal evolution. Finally, given the results of our field experiments indicating sequential assessment of divergent song and colour in the recognition of conspecifics, we discuss the consequences of multimodal signal divergence for the origin of species, as changes in signals across different sensory modalities may influence the evolution of premating reproductive isolation.     

Nitrate storage by phytoplankton in a coastal upwelling environment

The storage of nitrate by phytoplankton cells during the early phases of upwelling was studied in coastal stations off northern Spain (southern Bay of Biscay) between 1990 and 1994. In this region, a persistent upwelling during summer is characterised by intermittent pulses of variable intensity, and increased nutrient concentrations in the surface layer. The main effect of an upwelling pulse on phytoplankton distribution is the shifting of the chlorophyll a and primary production maxima to near the surface. When the upwelling relaxes, thermal stratification of the water column occurs, and a distinct subsurface chlorophyll maximum develops below the production maximum. An accumulation of intracellular nitrate characterized the early phases of upwelling (mean = 2.73 μmol N m⁻³), maximum concentrations being attained at depths where biomass and production values were moderate. In contrast, phytoplankton cells from non-upwelling situations contained significantly lower concentrations of intracellular nitrate (mean = 0.17 μmol N m⁻³). The variations in the intracellular pool of nitrate may result from the differential allocation of resources within the cell as a result of variations in the energy available, since the uptake and assimilation of nitrate is a relatively expensive process involving several enzymatic systems. We hypothesize that nitrate storage by phytoplankton cells is characteristic of early phases of upwelling and is linked to patterns of carbon fixation. Average nitrogen budgets for upwelling and non-upwelling situations indicate that intracellular nitrate reserves are not responsible for maintaining high phytoplankton growth rates, since they only account for <2% of daily primary production during upwelling events. Received: 28 August 1996 / Accepted 3 December 1996     

Variation in litter size: a test of hypotheses in Richardson's ground squirrels

We studied litter size variation in a population of Richardson's ground squirrels (Spermophilus richardsonii) in Alberta, Canada, from 1987 to 2004. Litter size at first emergence of juveniles from the natal burrow ranged from 1 to 14; the most common litter sizes, collectively accounting for 41.0% of 999 litters, were 6 and 7. The number of offspring surviving to adulthood (attained on emergence from hibernation as yearlings) increased with increasing litter size, a result that was not predicted by Lack's "optimal litter size" hypothesis, Mountford's "cliff-edge" effect, or the "bad-years" effect. Contrary to the negative effects predicted by the "cost of reproduction" hypothesis, litter size had no significant influence on survival of mothers to the subsequent year or on the size of the subsequent litter. Rather, our results best fit the predictions of the "individual optimization" hypothesis, which suggests that litter size is determined by the body condition and environmental circumstances of each mother. Supporting this hypothesis, survival of individual offspring was not significantly associated with litter size. Additionally, year-to-year changes in maternal body mass at mating were positively associated with concurrent changes in litter size (r = 0.56), suggesting that litter size depends on the body condition of the mother. Because the mean number of recruits to adulthood increased as litter size increased (r2 = 0.96) and litter size increased with maternal condition, offspring productivity was greater for mothers in better body condition.     

磷输入对沼泽湿地小叶章枯落物分解过程酶活性的影响

在三江平原沼泽湿地布置外源磷输入小区,选取典型湿地植物小叶章Calamagrostis angustifolia枯落物为研究对象,采用分解袋法,研究了不同梯度外源磷输入(CK:0;P1:1.2 g·m-2·a-1;P2:4.8 g·m-2·a-1;P3:9.6 g·m-2·a-1)对小叶章枯落物分解过程中酶活性的影响.结果表明,在小叶章枯落物分解前期,低磷输入(P1)抑制了蔗糖酶和脲酶活性,继续分解的过程中则极显著促进了二者的酶活性(P<0.01),而低磷输入对淀粉酶活性的影响则与之相反;中磷输入(P2)在小叶章枯落物分解的整个过程都促进了蔗糖酶和淀粉酶的活性,对脲酶活性除在分解65 d时有抑制,其他时期均提高了其酶活性;高磷输入(P3)均提高了枯落物分解后期的蔗糖酶和脲酶活性,其对淀粉酶活性的影响无明显规律;3种磷输入均显著抑制了酸性磷酸酶活性(P<0.01).     

Delayed maturation of multiple signals in a migratory songbird

Many animals use multiple signals in sexual communication, but our understanding of the interactions between multiple signals, particularly in inexperienced breeders, is limited. In birds, delayed plumage maturation (DPM) is well documented; young birds appear duller than adults, despite reaching sexual maturity. Age-related changes in song structure are also common in songbirds, though the extent to which songs of yearling males differ from those of adults (delayed song maturation, DSM) and its prevalence in species with DPM is unknown. We tested for DSM in the mate-attraction (repeat) song of a species with dramatic DPM, the American redstart (Setophaga ruticilla). Repeat song structure of territorial yearling and adult males differed significantly, based on discriminant analysis of nine non-collinear song features. Combined with previous evidence of delayed maturation in territorial (serial) song, we provide the first evidence for DSM in different song types used in different behavioral contexts during the breeding season of a migratory songbird. Within adults, variation in repeat song was associated with pairing, earlier onset of breeding, and number of offspring sired, suggesting a potential benefit for expressing more adult-like song. We found no relationship between the expression of adult-like repeat song and plumage in either age class. These results indicate that delayed maturation of repeat song in yearling redstarts is not due to a generalized delay in signal development, and suggest further work is needed to determine whether mate attraction (for both yearling and adult males) provides greater pressure to sound more “adult-like” than to look more “adult-like”.     

Prey capture and structure of the visual space of an insect that hunts by sight on the litter layer (Notiophilus biguttatus F., Carabidae,Coleoptera)

Summary The carabid beetle Notiophilus biguttatus hunts springtails and mites by visual cues. The preycapture behaviour of the beetle and the escape behaviour of the springtails were analysed by means of highspeed films. N. biguttatus has between 900 and 1250 ommatidia in each compound eye. The visual space covers ca. 200° in the horizontal plane, with a binocular overlap of no more than 74°. The fovea, the part of the eye where the pseudopupil is largest, points straight ahead of a beetle in its normal posture.The structure of the visual space was determined from measurements of the optical axes in the horizontal plane (plane of fixation) over the middle of the eye. Because of the slanted position of the ommatidia under the cornea, the optical axes point more towards the front or the back of the animal than do the anatomical axes.The optical axes were used to construct the binocular visual space in the horizontal plane. The point E , to which an estimation of distance is possible, lies on the midline 42.6 mm away from the front edges of the eyes. Resolution rapidly decreases with increasing distance, particularly depth resolution.At a distance corresponding to that from which the beetle attacks its prey, depth and width resolution correspond roughly to the dimensions of the smallest prey animals. The smallest measured directional corrections made by the beetle prior to attack (2°–3°) correspond approximately to the divergence angles in the fovea ( _h=2.2°), and the smallest measured distance correction prior to attack (0.2 mm) corresponds approximately to the depth resolution at attack distance.Supported by the Deutsche Forschungsgemeinschaft (SFB 4)     

Food-exchange by foragers in the hive – a means of communication among honey bees?

Dancing and trophallactic behaviour of forager honey bees, Apis mellifera ligustica >Spinola, that returned from an automatic feeder with a regulated flow rate of 50% weight-to-weight sucrose solution (range: 0.76–7.65 μl/min) were studied in an observation hive. Behavioural parameters of dancing, such as probability, duration and dance tempo, increased with the nectar flow rate, though with very different response curves among bees. For trophallaxis (i.e. mouth-to-mouth exchange of food), the frequency of giving-contacts and the transfer rate of the nectar increased with the nectar flow rate. After unloading, foragers often approached other nest mates and begged for food before returning to the food source. This behaviour was less frequent at higher nectar flow rates. These results show that the profitability of a food source in terms of nectar flow rate had a quantitative representation in the hive through quantitative changes in trophallactic and dancing behaviour. The role of trophallaxis as a communication channel during recruitment is discussed. Received: 14 January 1995/Accepted after revision: 14 August 1995     

Nonadditive effects of leaf litter species diversity on breakdown dynamics in a detritus-based stream

Since species loss is predicted to be nonrandom, it is important to understand the manner in which those species that we anticipate losing interact with other species to affect ecosystem function. We tested whether litter species diversity, measured as richness and composition, affects breakdown dynamics in a detritus-based stream. Using full-factorial analyses of single- and mixed-species leaf packs (15 possible combinations of four dominant litter species; red maple [Acer rubrum], tulip poplar [Liriodendron tulipifera], chestnut oak [Quercus prinus], and rhododendron [Rhododendron maximum]), we tested for single-species presence/absence (additive) or species interaction (nonadditive) effects on leaf pack breakdown rates, changes in litter chemistry, and microbial and macroinvertebrate biomass. Overall, we found significant nonadditive effects of litter species diversity on leaf pack breakdown rates, which were explained both by richness and composition. Leaf packs containing higher litter species richness had faster breakdown rates, and antagonistic effects of litter species composition were observed when any two or three of the four litter species were mixed. Less-consistent results were obtained with respect to changes in litter chemistry and microbial and macroinvertebrate biomass. Our results suggest that loss of litter species diversity will decrease species interactions involved in regulating ecosystem function. To that end, loss of species such as eastern hemlock (Tsuga canadensis) accompanied by predicted changes in riparian tree species composition in the southeastern United States could have nonadditive effects on litter breakdown at the landscape scale.